暗间断延迟玉米开花的相关分子机理研究
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摘要
玉米起源于热带南美洲的墨西哥地区,短日的光周期条件才能诱导正常开花。虽然经过了长期的人工选择和自然进化,大多数温带玉米对光周期不敏感,但是多数热带玉米在长日照光周期条件下,仍然表现出光周期敏感现象。光周期敏感是温带玉米育种中利用热带种质资源的主要限制因素。所以,解析光周期敏感的机理对于玉米遗传育种研究就显得非常重要。在短日植物的光周期诱导成花中,暗期是成花的决定因素,暗间断是在短日植物生长发育的光周期敏感阶段,于暗期的中间短暂曝光,这种处理往往会抑制或延迟短日植物开花。因此,利用暗间断处理也是研究玉米光周期敏感的一个重要方法。本研究利用暗间断处理的方法,从表型和光周期途径信号传导的上、中、下游关键基因表达规律的变化等角度,试图解析玉米光周期敏感的分子机理。本研究首先以热带和温带玉米自交系为材料,分析了暗间断处理后玉米生长发育的特点；其次,克隆了玉米光周期途径上游光信号受体光敏色素基因phyB2,通过荧光定量PCR分析该基因在暗间断处理前后48小时的表达规律；最后,研究暗间断处理后光周期途径中控制生物钟的两个关键基因Hd6、 ZmCCA1以及信号输出关键基因GI-CO-FT的表达规律及其与玉米开花的关系。得出如下儿个研究结果：
1、将热带玉米自交系CML288和温带玉米自交系黄早四与B73种植在短日条件下(9小时光/5小时暗),在光周期敏感时期进行暗间断,观测抽雄期、散粉期、吐丝期以及株高等的变化。结果表明,短日条件下连续暗间断处理后CML228抽雄期、散粉期、吐丝期推迟,株高降低；黄早四抽雄期、散粉期、吐丝期提前,株高增加；B73抽雄期、散粉期、吐丝期稍微推迟,株高略有降低。说明不同自交系对暗间断处理的反应不同,热带光周期敏感自交系对暗间断处理最敏感。对4-8叶期的不同发育时期暗间断处理进行比较发现,热带敏感自交系CML288在4叶期暗间断处理与短日照下的花期等性状没有差异,而在5叶期暗间断处理后抽雄期、散粉期、吐丝期推迟的天数最多,株高降低最大,说明5叶期是暗间断敏感的关键时期。在光周期敏感的5-8叶期之间,随着暗间断处理天数的增加,抽雄期、散粉期、吐丝期推迟的天数越多,说明暗间断处理具有一定的累加效应。用白光、红光和远红光进行暗间断,发现红光的效应最大。红光的暗间断效应可以被随后的远红光所逆转,表明光敏素是玉米主要的响应于暗间断的光接收器。
2、利用同源克隆技术,成功地克隆了phyB2的CDS序列。CDS序列全长3501bp,编码一条1166个氨基酸残基的肽链,包含6个保守结构域。黄早四与CML288的phyB2基因cDNA序列对比发现,一共存在17处单碱基的变异。其中,7处单碱基的变异引起了编码氨基酸的变化,而且有3处差异位于PHYB2保守的结构域内。研究了phyB2在短日照和暗间断后48小时内的表达节律。在短日照条件下,CML288叶片和茎尖中phyB2呈现双峰的表达模式,其表达峰值出现在暗期结束时和进入暗期后第6时；暗间断处理后,phyB2也呈现双峰的表达模式,但是其表达峰值比短日照条件下提前。在短日照条件下,黄早4叶片和茎尖中phyB2呈现双峰的表达模式,其表达峰值出现在暗期结束前的第3小时和暗期开始时;暗间断处理后,phyB2的表达模式基本没有改变。
3、研究了自交系CML288和黄早四中生物钟基因Hd6、ZmCCA124小时在叶片和茎尖中短日照和暗间断处理后的表达规律。结果表明,在短日照条件下,Hd6基因在CML288的叶片和茎尖中光期表达量较低,进入暗期后在叶片中开始上升,在茎尖中光期前第4小时出现表达峰值；暗间断处理后,Hd6的表达量有所降低。在短日条件下,ZmCCA1基因在CML288叶片和茎尖中表达峰值都是出现在光期后第4小时。暗间断提高了ZmCCA1基因mRNA的积累,但是并没有改变这个基因在短日条件下的表达节律。
短日照条件下,Hd6基因在黄早四的叶片和茎尖中的表达量从光期后第4小时开始升高,在叶片中暗期后第3小时达到表达峰值,而在茎尖中光期前第4小时达到表达峰值。暗间断处理后,Hd6基因在黄早四的叶片和茎尖中表达量降低,进入光期后很快就恢复为止常短日条件下的表达规律。短日条件下,ZmCCA1基因在黄早四叶片和茎尖中表达规律和CML288类似。暗间断处理提高了黄早四中ZmCCA1基因在暗期的表达量,但是在进入光期以后,ZmCCA1基因的表达就近于正常。
4、CML288在短日照条件下,叶片和茎尖中的ZmGI在暗期前第3小时达到峰值；暗间断处理没有改变叶片中ZmGI的表达节律,却提高了它的表达量,但茎尖中ZmGI的表达峰值提前。短日条件下,CML288叶片和茎尖中conz1基因表达峰值出现在暗期后第6小时；暗间断使叶片中的conz1基因表达节律呈现双峰模式,而茎尖中conz1基因的表达水平一直处于较低状态,进入暗期后第3小时达到峰值。在短日条件下,ZCN8基因在CML288叶片和茎尖中暗期结束时表达量最高,夜晚表达量低；暗间断强烈抑制了ZCN8基因在CML288叶片和茎尖中的表达水平,在下一个周期表达量才开始恢复。
在短日条件下,ZmGI基因在黄早四叶片和茎尖中的表达峰值出现在暗期前第3小时；暗间断处理后,推迟了叶片中ZmGI表达峰值出现的时间,提高了表达峰值,没有改变茎尖中ZmGI原有的表达节律,而是峰值有所降低。短日和暗间断条件下,conz1基因在黄早四叶片和茎尖中的表达规律基本一致。在短日条件下,ZCN8基因在黄早四中暗期结束时达到峰值,暗间断降低了ZCN8的表达水平,但是没有改变ZCN8的固有表达节律。
5、暗间断处理光周期敏感的玉米自交系CML288后,phyB2基因峰值提前,可能使生物钟基因Hd6表达量降低和CCA1表达量升高,延长了生物钟周期,使生物钟信号输出基因GI表达量升高,进而引起conz1基因的表达类似于长日下的双峰表达模式；同时,暗间断处理也可能引起conz1基因出现双峰表达模式,从而抑制ZCN8基因的表达,最终引起开花延迟。
ABSTRACT:Maize (Zea mays L.) is domesticated from Mexico of south American, a plant requiring short day photoperiods to flower. Although the majority of maize varieties are generally characterized as day-neutral plants through centuries of domestication and artificial selection, numerous tropical landraces of maize remain sensitive to photoperiod. Photoperiod sensitivity was the major obstacle to using tropical for breeding programs of temperate maize. Therefore, it is vital for maize genetics and breeding to understand the genetic basis of photoperiod sensitivity (PS).Dark period is the main facter in flowering induce of short-day plants (SDPs). A short exposure to light in the middle of the night causes inhibition of flowering in SDPs, this phenomenon is called night break (NB) and has been used as a tool to study the phothperiodic control of flowering. So, NB is also an important method to study maize PS.
In this study, the genetic basis of maize photoperiod sensitivity were carried out from the aspects of phenotype and photoperiod signaling passway by using NB. Firstly, tropical and temperate maize inbred lines were conducted by NB to analyse characters of development traits. Secondly, phyB2as a main photoreceptor of maize were cloned, and their expression patterns were analysed before and after NB within48h by real-time RT-PCR. Finally, expression of circadian clock genes Hd6, ZmCCA1and signaling output pathway genes Gigantea (GI), Constans(CO), and Flowering Locus T (FT) were addressed to investigate the relationship between their expression patterns and maize flowering. The main results obtained-in this research were as following:
1. A tropical maize inbred line CML288and two temperate inbred line B73, Huangzaosi were grown under inductive short-day (SD)9h light/15h dark cycles, continuous night break (NB) under SD condition were conducted in the most photoperiod sensitive developmental stage. Changes in day to tassel (DTT), day to pollen (DTP), day to silking (DTS) and plant height were measured after NB. The results indicated that DTT, DTP, DTS were delayed, and the plant height was decreased in CML288after NB, so is in B73with lower degree, while DTT, DTP, DTS were advanced, and the plant height was increased in Huangzaisi. All these demonstrated that different lines produce different NB responses and tropical photoperiod sensitive inbred line is the most sensitive to NB. NB effects were compared from4-leaf period to8-leaf period, the results showed that traits in flowering time and plant height of tropical photoperiod sensitive inbred line CML288were similar to that of SD condition when NB conducted at4-leaf period. The largest effects of NB on these traits occurred when NB were conducted at5th "leaf period, demonstrating that the5th-leaf stage is the most sensitive to NB. The dalay of DTT, DTP, DTS and inhibition in plant height were positively correlated to the days of NB treatment when continuous NB were conducted at the photoperiod sensitive developmental5-leaf to8-leaf period, indicated that NB treatment has additive effects.Red light(R) and Far Red light(FR) irradiation in the middle of night period also delayed flowering time of CML288,effect of R on inhibition of flowering was larger than that of FR, and FR could partly reverse these effect.These results implied that phytochrome was the photoreceptor in the NB effects on growth and development of maize.
2.CDS of phyB2was cloned by homologous amplification in maize inbred lines CML288and Huangzaisi, a3501bp open reading encoded1166amino acids which has six conserved domains.There are seventeen single DNA letter differences in phyB2CDS of CML288and Huangzaisi. Multiple alignments of the predicted complete amino acid sequences of these two lines indicated that there are seven amino acid diversity, three of seven were located in conserved domains of PHYB2. Expression pattern of phyB2in48h under SD and NB conditions were conducted using real-time fluorescence quantitative PCR in the stem apexes and leaves of CML288and Huangzaisi. Under SD conditions, phyB2exprssion patterns in leaves and SAs of CML288peaked twice, the first peak occurred at dawn,the second peak occurred at6h after dusk. NB treatment brought forward the peak of phyB2in leaves of CML288. phyB2exprssion patterns in leaves and SAs of Huangzaisi.also peaked twice under SD conditions, the peaks occurred at3h before dawn and dusk, expression patterns hardly changed after NB.
3.Expression pattern of circadian clock genes Hd6、ZmCCAl in24h under SD and NB conditions were conducted in the stem apexes and leaves of CML288and-Huangzaisi. Under SD conditions, Hd6exprssion levels was lower in leaves and SAs during light period, then increased in leaves after entering dark period, while peaked at4h before dawn. Hd6exprssion levels decreased after NB treatment. Under SD conditions, ZmCCAl exprssion levels peaked at at4h after dawn. NB treatment did not change this gene exprssion pattern but enhanced its level.
Under SD conditions, Hd6exprssion levels in leaves and SAs of Huangzaisi increased at4h after dawn, then peaked at3h after dusk in leaves, while peaked at4h before dawn in SAs. NB treatment decreased its exprssion levels in leaves and SAs, then recovered to normal SD expression pattern after dawn. ZmCCAI exprssion patterns in leaves and SAs of Huangzaisi was similar to CML288, NB treatment increased its exprssion levels during dark period, but recovered to normal level after dawn.
4.Under SD conditions, ZmGI exprssion peaked at3h before dawn in leaves and SAs of CML288. NB treatment did not change its exprssion pattern, but enhanced its exprssion levels in leaves, while brought forward its peak in SAs. conzl exprssion levels in leaves and SAs peaked at6h after dusk under SD conditions, conzl expression in leaves exhibited a diurnal expression pattern.with two peaks per day while its levels in SAs were lower all the time by NB treatment. ZCN8exprssion levels in leaves and SAs peaked at dawn under SD conditions, NB treatment dramatically inhibited its levels,but recovered to normal at the second cycle.
Under SD conditions, ZmGI exprssion levels in leaves and SAs of Huangzaisi peaked at3h before dusk. NB treatment delayed its peak and enhanced its amplitude in leaves,while decreased its amplitude in SAs with normal expression pattern, conzl exprssion patterns in leaves and SAs was similar to that under SD conditions and NB treatment. ZCN8exprssion levels peaked at dawn under SD conditions. NB treatment decreased its amplitude,but did not changed its normal expression pattern.
5.After NB treatment, phyB2peaked earlier in tropical photoperiod sensitive inbred line CML288,this result decreased circadian clock genes Hd6level, and increased ZmCCA1level,giving rise to longer circadian clock period. Longer circadian clock period induced higher ZmGI exprssion levels to produce two peaks expression pattern of conzl gene, inhibit ZCN8exprssion, resulting in flowering delay in the end.
1、将热带玉米自交系CML288和温带玉米自交系黄早四与B73种植在短日条件下(9小时光/5小时暗),在光周期敏感时期进行暗间断,观测抽雄期、散粉期、吐丝期以及株高等的变化。结果表明,短日条件下连续暗间断处理后CML228抽雄期、散粉期、吐丝期推迟,株高降低；黄早四抽雄期、散粉期、吐丝期提前,株高增加；B73抽雄期、散粉期、吐丝期稍微推迟,株高略有降低。说明不同自交系对暗间断处理的反应不同,热带光周期敏感自交系对暗间断处理最敏感。对4-8叶期的不同发育时期暗间断处理进行比较发现,热带敏感自交系CML288在4叶期暗间断处理与短日照下的花期等性状没有差异,而在5叶期暗间断处理后抽雄期、散粉期、吐丝期推迟的天数最多,株高降低最大,说明5叶期是暗间断敏感的关键时期。在光周期敏感的5-8叶期之间,随着暗间断处理天数的增加,抽雄期、散粉期、吐丝期推迟的天数越多,说明暗间断处理具有一定的累加效应。用白光、红光和远红光进行暗间断,发现红光的效应最大。红光的暗间断效应可以被随后的远红光所逆转,表明光敏素是玉米主要的响应于暗间断的光接收器。
2、利用同源克隆技术,成功地克隆了phyB2的CDS序列。CDS序列全长3501bp,编码一条1166个氨基酸残基的肽链,包含6个保守结构域。黄早四与CML288的phyB2基因cDNA序列对比发现,一共存在17处单碱基的变异。其中,7处单碱基的变异引起了编码氨基酸的变化,而且有3处差异位于PHYB2保守的结构域内。研究了phyB2在短日照和暗间断后48小时内的表达节律。在短日照条件下,CML288叶片和茎尖中phyB2呈现双峰的表达模式,其表达峰值出现在暗期结束时和进入暗期后第6时；暗间断处理后,phyB2也呈现双峰的表达模式,但是其表达峰值比短日照条件下提前。在短日照条件下,黄早4叶片和茎尖中phyB2呈现双峰的表达模式,其表达峰值出现在暗期结束前的第3小时和暗期开始时;暗间断处理后,phyB2的表达模式基本没有改变。
3、研究了自交系CML288和黄早四中生物钟基因Hd6、ZmCCA124小时在叶片和茎尖中短日照和暗间断处理后的表达规律。结果表明,在短日照条件下,Hd6基因在CML288的叶片和茎尖中光期表达量较低,进入暗期后在叶片中开始上升,在茎尖中光期前第4小时出现表达峰值；暗间断处理后,Hd6的表达量有所降低。在短日条件下,ZmCCA1基因在CML288叶片和茎尖中表达峰值都是出现在光期后第4小时。暗间断提高了ZmCCA1基因mRNA的积累,但是并没有改变这个基因在短日条件下的表达节律。
短日照条件下,Hd6基因在黄早四的叶片和茎尖中的表达量从光期后第4小时开始升高,在叶片中暗期后第3小时达到表达峰值,而在茎尖中光期前第4小时达到表达峰值。暗间断处理后,Hd6基因在黄早四的叶片和茎尖中表达量降低,进入光期后很快就恢复为止常短日条件下的表达规律。短日条件下,ZmCCA1基因在黄早四叶片和茎尖中表达规律和CML288类似。暗间断处理提高了黄早四中ZmCCA1基因在暗期的表达量,但是在进入光期以后,ZmCCA1基因的表达就近于正常。
4、CML288在短日照条件下,叶片和茎尖中的ZmGI在暗期前第3小时达到峰值；暗间断处理没有改变叶片中ZmGI的表达节律,却提高了它的表达量,但茎尖中ZmGI的表达峰值提前。短日条件下,CML288叶片和茎尖中conz1基因表达峰值出现在暗期后第6小时；暗间断使叶片中的conz1基因表达节律呈现双峰模式,而茎尖中conz1基因的表达水平一直处于较低状态,进入暗期后第3小时达到峰值。在短日条件下,ZCN8基因在CML288叶片和茎尖中暗期结束时表达量最高,夜晚表达量低；暗间断强烈抑制了ZCN8基因在CML288叶片和茎尖中的表达水平,在下一个周期表达量才开始恢复。
在短日条件下,ZmGI基因在黄早四叶片和茎尖中的表达峰值出现在暗期前第3小时；暗间断处理后,推迟了叶片中ZmGI表达峰值出现的时间,提高了表达峰值,没有改变茎尖中ZmGI原有的表达节律,而是峰值有所降低。短日和暗间断条件下,conz1基因在黄早四叶片和茎尖中的表达规律基本一致。在短日条件下,ZCN8基因在黄早四中暗期结束时达到峰值,暗间断降低了ZCN8的表达水平,但是没有改变ZCN8的固有表达节律。
5、暗间断处理光周期敏感的玉米自交系CML288后,phyB2基因峰值提前,可能使生物钟基因Hd6表达量降低和CCA1表达量升高,延长了生物钟周期,使生物钟信号输出基因GI表达量升高,进而引起conz1基因的表达类似于长日下的双峰表达模式；同时,暗间断处理也可能引起conz1基因出现双峰表达模式,从而抑制ZCN8基因的表达,最终引起开花延迟。
ABSTRACT:Maize (Zea mays L.) is domesticated from Mexico of south American, a plant requiring short day photoperiods to flower. Although the majority of maize varieties are generally characterized as day-neutral plants through centuries of domestication and artificial selection, numerous tropical landraces of maize remain sensitive to photoperiod. Photoperiod sensitivity was the major obstacle to using tropical for breeding programs of temperate maize. Therefore, it is vital for maize genetics and breeding to understand the genetic basis of photoperiod sensitivity (PS).Dark period is the main facter in flowering induce of short-day plants (SDPs). A short exposure to light in the middle of the night causes inhibition of flowering in SDPs, this phenomenon is called night break (NB) and has been used as a tool to study the phothperiodic control of flowering. So, NB is also an important method to study maize PS.
In this study, the genetic basis of maize photoperiod sensitivity were carried out from the aspects of phenotype and photoperiod signaling passway by using NB. Firstly, tropical and temperate maize inbred lines were conducted by NB to analyse characters of development traits. Secondly, phyB2as a main photoreceptor of maize were cloned, and their expression patterns were analysed before and after NB within48h by real-time RT-PCR. Finally, expression of circadian clock genes Hd6, ZmCCA1and signaling output pathway genes Gigantea (GI), Constans(CO), and Flowering Locus T (FT) were addressed to investigate the relationship between their expression patterns and maize flowering. The main results obtained-in this research were as following:
1. A tropical maize inbred line CML288and two temperate inbred line B73, Huangzaosi were grown under inductive short-day (SD)9h light/15h dark cycles, continuous night break (NB) under SD condition were conducted in the most photoperiod sensitive developmental stage. Changes in day to tassel (DTT), day to pollen (DTP), day to silking (DTS) and plant height were measured after NB. The results indicated that DTT, DTP, DTS were delayed, and the plant height was decreased in CML288after NB, so is in B73with lower degree, while DTT, DTP, DTS were advanced, and the plant height was increased in Huangzaisi. All these demonstrated that different lines produce different NB responses and tropical photoperiod sensitive inbred line is the most sensitive to NB. NB effects were compared from4-leaf period to8-leaf period, the results showed that traits in flowering time and plant height of tropical photoperiod sensitive inbred line CML288were similar to that of SD condition when NB conducted at4-leaf period. The largest effects of NB on these traits occurred when NB were conducted at5th "leaf period, demonstrating that the5th-leaf stage is the most sensitive to NB. The dalay of DTT, DTP, DTS and inhibition in plant height were positively correlated to the days of NB treatment when continuous NB were conducted at the photoperiod sensitive developmental5-leaf to8-leaf period, indicated that NB treatment has additive effects.Red light(R) and Far Red light(FR) irradiation in the middle of night period also delayed flowering time of CML288,effect of R on inhibition of flowering was larger than that of FR, and FR could partly reverse these effect.These results implied that phytochrome was the photoreceptor in the NB effects on growth and development of maize.
2.CDS of phyB2was cloned by homologous amplification in maize inbred lines CML288and Huangzaisi, a3501bp open reading encoded1166amino acids which has six conserved domains.There are seventeen single DNA letter differences in phyB2CDS of CML288and Huangzaisi. Multiple alignments of the predicted complete amino acid sequences of these two lines indicated that there are seven amino acid diversity, three of seven were located in conserved domains of PHYB2. Expression pattern of phyB2in48h under SD and NB conditions were conducted using real-time fluorescence quantitative PCR in the stem apexes and leaves of CML288and Huangzaisi. Under SD conditions, phyB2exprssion patterns in leaves and SAs of CML288peaked twice, the first peak occurred at dawn,the second peak occurred at6h after dusk. NB treatment brought forward the peak of phyB2in leaves of CML288. phyB2exprssion patterns in leaves and SAs of Huangzaisi.also peaked twice under SD conditions, the peaks occurred at3h before dawn and dusk, expression patterns hardly changed after NB.
3.Expression pattern of circadian clock genes Hd6、ZmCCAl in24h under SD and NB conditions were conducted in the stem apexes and leaves of CML288and-Huangzaisi. Under SD conditions, Hd6exprssion levels was lower in leaves and SAs during light period, then increased in leaves after entering dark period, while peaked at4h before dawn. Hd6exprssion levels decreased after NB treatment. Under SD conditions, ZmCCAl exprssion levels peaked at at4h after dawn. NB treatment did not change this gene exprssion pattern but enhanced its level.
Under SD conditions, Hd6exprssion levels in leaves and SAs of Huangzaisi increased at4h after dawn, then peaked at3h after dusk in leaves, while peaked at4h before dawn in SAs. NB treatment decreased its exprssion levels in leaves and SAs, then recovered to normal SD expression pattern after dawn. ZmCCAI exprssion patterns in leaves and SAs of Huangzaisi was similar to CML288, NB treatment increased its exprssion levels during dark period, but recovered to normal level after dawn.
4.Under SD conditions, ZmGI exprssion peaked at3h before dawn in leaves and SAs of CML288. NB treatment did not change its exprssion pattern, but enhanced its exprssion levels in leaves, while brought forward its peak in SAs. conzl exprssion levels in leaves and SAs peaked at6h after dusk under SD conditions, conzl expression in leaves exhibited a diurnal expression pattern.with two peaks per day while its levels in SAs were lower all the time by NB treatment. ZCN8exprssion levels in leaves and SAs peaked at dawn under SD conditions, NB treatment dramatically inhibited its levels,but recovered to normal at the second cycle.
Under SD conditions, ZmGI exprssion levels in leaves and SAs of Huangzaisi peaked at3h before dusk. NB treatment delayed its peak and enhanced its amplitude in leaves,while decreased its amplitude in SAs with normal expression pattern, conzl exprssion patterns in leaves and SAs was similar to that under SD conditions and NB treatment. ZCN8exprssion levels peaked at dawn under SD conditions. NB treatment decreased its amplitude,but did not changed its normal expression pattern.
5.After NB treatment, phyB2peaked earlier in tropical photoperiod sensitive inbred line CML288,this result decreased circadian clock genes Hd6level, and increased ZmCCA1level,giving rise to longer circadian clock period. Longer circadian clock period induced higher ZmGI exprssion levels to produce two peaks expression pattern of conzl gene, inhibit ZCN8exprssion, resulting in flowering delay in the end.
引文
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